Deep Coral and Associated Species Taxonomy and Ecology Deepcast II Expedition Report

Deep Coral and Associated Species Taxonomy and Ecology DeepCAST II Expedition Report

Roatán, Honduras. May 21‐28, 2011.

Report date: September 2011

NOAA Technical Memorandum NOS NCCOS 137

This report has been reviewed by the National Ocean Service of the National Oceanic and Atmospheric Administration (NOAA) and approved for publication. Such approval does not signify that the contents necessarily represent the official position of NOAA or of the Government of the United States, nor does mention of trade names or commercial products constitute endorsement or recommendation for their use. Furthermore, this report is intended to serve as a record of cruise activities and preliminary observations during a field survey in support of an ongoing research study, and thus does not represent a final analysis or interpretation of project results.

Cover Photo. A large yellow sea fan colony with Asteroschema sp. brittlestars in the branches and an orange colored brisingid seastar Novodinia antillensis on a rocky outcrop at 700 meters depth in Roatán, Honduras.

Citation for this Report: Etnoyer, PJ, TC Shirley, KA Lavelle. 2011. Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) II Expedition Report. NOAA Technical Memorandum NOS NCCOS 137. NOAA/NOS Center for Coastal Environmental Health and Biomolecular Research, Charleston, SC. 42 pp.

Expedition Report Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) II Expedition, May 21‐28, 2011

Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) II Expedition Report

Peter J. Etnoyer1, Thomas C. Shirley2, and Katherine A. Lavelle2

1 NOAA Ocean Service, National Centers for Coastal Ocean Science (NCCOS), Center for Coastal Environmental Health and Biomolecular Research (CCEHBR), 219 Fort Johnson Road, Charleston, SC, 29412‐9110

2 Texas A&M University‐ Corpus Christi, 6300 Ocean Drive, Corpus Christi, TX, 78412

[Also see list of other collaborating scientists on the final page of this report]

NOAA Technical Memorandum NOS NCCOS 137

September 2011

United States Department of National Oceanic and National Ocean Service Commerce Atmospheric Administration

Gary Locke Jane Lubchenco David Kennedy (A) Secretary Administrator Assistant Administrator

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Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) II Expedition Roatán, Honduras. May 21-28, 2011.

Introduction:

NOAA has a mandate to explore and understand deep-sea coral ecology under Magnuson-Stevens Sustainable Fisheries Conservation Act Reauthorization of 2009. Deep-sea corals are increasingly considered a proxy for marine biodiversity in the deep-sea because corals create complex structure, and this structure forms important habitat for associated species of shrimp, crabs, sea stars, brittle stars, and fishes. Yet, our understanding of the nature of the relationships between deep-corals and their associated species is incomplete.

One of the primary challenges of conducting any type of deep-sea coral (DSC) research is access to the deep-sea. The deep-sea is a remote environment that often requires long surface transits and sophisticated research vehicles like submersibles and remotely operated vehicles (ROVs). The research vehicles often require substantial crew, and the vehicles are typically launched from large research vessels costing many thousands of dollars a day.

To overcome the problem of access to the deep-sea, the Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) Expeditions are pioneering the use of shore-based submersibles equipped to do scientific research. Shore-based subs alleviate the need for expensive ships because they launch and return under their own power. One disadvantage to the approach is that shore-based subs are restricted to nearby sites. The disadvantage is outweighed, however, by the benefit of repeated observations, and the opportunity to reduce the costs of exploration while expanding knowledge of deep-sea coral ecology.

Expedition Summary:

The DeepCAST II Expedition explored depths below the Meso-American Reef in Roatán, Honduras from 21- 28 May 2011. The team explored deep-sea habitats using the Roatán Institute for Deep-sea Exploration submersible Idabel, captained by Karl Stanley. The team made six dives between 1200 and 2200 feet deep (365 and 670 meters), conducting photo and video transects to estimate coral and sponge diversity and abundance, characterizing water chemistry, and collecting corals and mollusks to discern the relationships between corals and their epifauna.

Among the highlights of the trip were:

 First confirmed documentation of live Lophelia pertusa along the Meso-American Reef.  Comprehensive sampling of slit shell diversity for genetic and biogeographic studies.  Live aggregations of very large, presumably very old, deep-sea gorgonian corals.  Octocoral diversity and biomass exceeding well-known sites in the Gulf of Mexico.  Numerous species in association with corals, including sea stars, sea urchins, brittle stars, mollusks, crabs, and fishes.  Evidence of agonistic interactions, including predation on deep-sea coral colonies.  Photographic data to measure coral recovery rates from prior predation events.  Evidence of human pollution (trash & fishing gear) degrading coral habitat near 1500 ft (460 m)

Project Description:

The Mesoamerican Reef (MAR) is the Atlantic Ocean’s largest coral reef, and the second largest barrier reef in the world. The reef stretches across 1000 km of the continental shelf from Honduras to Mexico (Fig. 1). The continental slope is directly adjacent to the reef, and drops abruptly to bathyal depths of 3500 m and more. The substrate and topography of the deep escarpment are well suited to deep-sea corals, including ‘stony’ reef building corals, soft corals, cup corals, sea fans, and black corals.

Yet, only 2 or 3 sites along the MAR have been explored using submersibles or sampled using trawls. Where the present study was conducted, rich and abundant assemblages of hard and soft corals were found. Species composition of the deep reef assemblage appears similar to neighboring US reefs, with many shared species. This suggests deep-sea communities along the MAR may be connected to the Gulf of Figure 1. Map of the Western Caribbean Sea showing the extent of Mexico and Straits of Florida through the deep Meso-American Reef, from Honduras (bottom) to larval dispersal and exchange. Yucatan Channel (top). The 800 m isobath is in red, showing the depth limit of our explorations. Isla de Roatán in the Bay Islands is located at the southern extent of the MAR, 50 km north of mainland Honduras. The island offers ready access to remarkable deep-sea coral aggregations 700 m and deeper. Roatán is home to the Idabel submersible and Roatán Institute for Deep-Sea Exploration (RIDE). RIDE was founded by Idabel designer, builder, and pilot Karl Stanley.

The Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) II Expedition used the Idabel submersible to collect photographic, video, and water chemistry data. The research team included marine biologists from National Oceanic and Atmospheric Administration (NOAA), Smithsonian Institution National Museum of Natural History (NMNH), Texas A&M University- Corpus Christi (TAMU-CC), and Conservation International (CI). Dr. Peter Etnoyer, NOAA, led the research expedition. Dr. Tom Shirley, TAMU-CC, and Dr. Jerry Harasewych, NMNH, served as co-leaders.

Dive Location:

Figure 2. Plan view of Roatán Island, situated 70 kilometers north of mainland Honduras. The pink dots indicate the location of six submersible dives conducted for DeepCAST II Expedition.

Project Goals and Objectives:

The primary goals of the DeepCAST expeditions were to: 1) estimate deep coral and sponge relative abundance, density, and diversity for comparison to other known aggregations in the Western Atlantic; and, 2) discern the nature of the relationships between host corals and their associated species, including epifauna living on the corals, and predators feeding on the corals. To accomplish these goals, the group conducted non-invasive photo and video transects of deep-coral fauna, deployed baited traps, and collected voucher specimens of corals (e.g., Lophelia pertusa and Dendrophyllia alternata) and associated species (e.g., gastropods and crinoids).

A secondary goal of the DeepCAST research was to characterize water column chemistry in and around Lophelia pertusa reefs. This was accomplished by mounting a Conductivity-Temperature-Depth- Dissolved Oxygen (CTD-O) with pH sensor to the Idabel submersible. CTD-O data were downloaded to a field laptop daily upon recovery. The data may help to define the “climate envelope” of Lophelia pertusa in the Caribbean Sea by providing information on temperature, salinity, oxygen, and pH. These parameters may be correlated to aragonite saturation state through future water sampling efforts.

Idabel submersible:

Idabel was used to gather biological samples, photo, video, CTD-O, and pH data. The sub is equipped with interior and exterior still/video cameras, synchronized exterior strobes, and parallel exterior lasers. The vessel is pressure rated to 1000 m depth by designer, builder, and pilot Karl Stanley. The vehicle has a weight limit of 450 lbs (204 kg) plus the pilot. Two scientists occupied the forward compartment.

Figure 3. Idabel submersible is a three-person sub with a 32in. (81 cm) viewport for scientific observers.

Dive Schedule: Table 1. DeepCAST II dive dates, observers, locations, depths, and activities. CTD-O = Conductivity-Temperature- Depth-Oxygen sensor, AKR = Anthony’s Key Resort.

Dive Max. Date Obs. 1 Obs. 2 Location Activity No. depth (m)

- 0 none none Half Moon Bay - deploy baited trap for biology 5/22/2011 1 Tom Kate Luna Beach 630 Photo-transect, CTD 5/23/2011 2 Jerry Sebastian Sueno del Mar 655 Photo-transect, CTD, samples

5/23/2011 3 Laney Annie Half Moon Bay 465 Video transect, CTD, recover trap AKR 1 - 5/25/2011 4 Kate Annie 660 Photo-transect, CTD, samples Three Kings AKR 2 - 5/26/2011 5 Sebastian Laney 640 Photo-transect, CTD, samples Lawson's Rock 5/27/2011 6 Tom Jerry Old Loafers 375 Photo-transect, CTD, samples

Photo and video sampling methodology:

The primary objective of DeepCAST II was to conduct quantitative photo and video transects of deep coral and sponge habitat in at least 4 independent sites in the 100-700 m depth range, using interior cameras synchronized to strobes mounted on the exterior of the submersible. Observers operated one camera continuously throughout the course of each 4-6 hour dive. The still camera operator took approximately 1 photo/minute, shooting wide and medium angle shots of the seafloor with particular attention to coral and sponge colonies. The video operator recorded continuous high-definition video on 1 hour-long Mini-DV tapes. Camera operators tried to frame the images to include the parallel green lasers projected from the sub onto the seafloor. All watches and cameras were synchronized with the CTD. Divers produced one-page summaries of each dive with dive chronologies providing information on dive start time, start coordinates, max depth, min depth, habitat, debris, and animals observed.

CTD-O sampling methodology:

A Seabird Seacat 19 conductivity-temperature-depth-oxygen (CTD-O) sensor with a supplementary pH sensor was mounted on the starboard side rail of Idabel submersible to generate continuous, along-track water chemistry profiles for each dive, especially around Lophelia pertusa reefs. The unit was activated before each dive, and data were downloaded on recovery. The unit did not record for the whole transit on most dives, but one complete track was recorded on Dive 6.

Biological sampling methodology:

Biological samples from the deep-sea were collected with a baited trap and a dip net mounted on the side of the Idabel submersible.

The following items were requested in a sampling permit application to Andrés Alegría, Coordinador de Monitoreo Biológico, Instituto Nacional de Conservación y Desarrollo Forestal, Áreas Protegidas y Vida Silvestre (ICF), Honduras.

Dr. Peter Etnoyer requested up to 5 specimens each of octocorals (sea fans) in the families Primnoidae, Ellisellidae, Isididae, and Paramuriceidae. Voucher specimens will be deposited in the collections of the Smithsonian Institution. If authorized, one or two additional specimens would be collected to be returned to the National University’s museum, as requested by Ian Drysdale.

Dr. Jerry Harasewych requested up to 3 specimens each of the deep-sea snails in the families Pleurotomariidae, Buccinidae, and Muricidae that are encountered during the dives. These would be deposited in the collection of the Smithsonian Institution as voucher specimens for anticipated publications. Additional specimens will be collected for the National Univ. museum.

Dr. Marcelo Kitahara could not participate, but requested 5 specimens of deep-water scleractinia Lophelia pertusa, Dendrophyllia alternata, Madrepora oculata, Javania cailleti, Desmophyllum dianthus and Caryophyllia spp. Voucher specimens will be deposited in the collections of the Smithsonian Institution. Additional specimens will be collected for the National Univ. museum.

Scientific apparatus:

Modifications to Idabel submersible for DeepCAST II included an exterior camera mount with parallel lasers for scale, mounted 10 cm apart, and a Seabird Seacat v 19 conductivity-temperature-depth-oxygen (CTD-O) sensor. Camera and lasers were mounted forward on the sub, below the viewport. The CTD-O was on a lower starboard side rail.

Figure 4. The laser mount (in black) is positioned below the camera housing (in yellow).

Figure 5. A Seabird Seacat 19 CTD-O water chemistry profiler mounted on the starboard rail.

Sampling apparatus:

Long-pole net: A long, yellow mesh net was weighted with stones to make it hang. The net was attached to a hardened steel hoop on the end of a 14ft (4.3 m) long steel pole. The pole was mounted to the port side rail of Idabel and used to collect voucher specimens of scleractinian corals (Fig. 6).

Figure 6. Lophelia pertusa A long-pole net is used to collect at 1920 ft (585 m) depth.

Baited trap: A green plastic basket was baited with lionfish heads, wrapped in mesh fabric, and weighted with rebar. Yellow rope was threaded through a block of buoyant syntactic foam and attached to the basket to create a loop for retrieval (Fig. 7). The retrieval operation used a long shepherd’s crook device mounted on the port side rail of Idabel. The crook successfully recovered the trap to the surface.

Figure 7. A sculptured lobster (Eunephrops cadenasi) and shrimp (Notostomus sp.) guard the baited trap at 1510 ft (460 m). Lasers are 10 cm, shown above, left of center. 7

Dive summaries:

DeepCAST II Dive 1 Date: May 22, 2011 Location: Luna Beach

Descent coordinates: 16° 17.691’ N, 86° 36.843’ W Observers: Tom Shirley, Kate Lavelle Left dock: 0941 hr Begin Descent: 1014 hr Return to dock: 1510 hr

Geology: Maximum depth of dive 2090 ft. (637 m) Many large basaltic boulders; one was 105 ft vertical relief. Boulders were interspersed with soft sediment. No patterns were observed between spacing of sediment between boulders; some distances between boulders were long, others were short. Many boulders supported little visible epifauna.

Biology: Soft substrate was occupied by sea urchins, shrimp, stalked crinoids, a few sponges, elasipodid sea cucumbers, and a few sharks. One cucumber was observed (and video-taped and photographed) swimming. Sea cucumbers, crinoids, and erect cnidarians colonies often tipped with sponges or anemones, were most common on soft substrate. Sea cucumbers were usually solitary, often several meters from other cucumbers. Stalked cnidarians colonies occurred in abundance, almost regularly spaced; stalked crinoids occurred in shallower depths (780 ft, 238 m) and also occurred in relatively dense assemblages. Shrimp occurred near some objects on the bottom, either a boulder or anemone. One aggregation of sea urchins, about 30 sea urchins, occurred on soft sediment.

Most common DSC on hard substrate were large Primnoidae, but in places yellow Paramuriceidae were abundant (at 13:44, 1110 ft, 338 m). DSC were usually most common on tops of boulders or on undersides of ledges along the steep slope. At 1400 ft. Parazooanthus coral occurred in dense clumps on tops of boulders in shallower water in a few discrete locations. Karl remarked that he had not seen these previously, but I believe that Peter and I observed these in August 2010. A new (for our observations) hemiuralyid brittlestar (Asteroporpa annulata) was observed on what appeared to be a yellow Paramuriceidae (but with slightly larger branches) in shallower water (850 ft).

A mass spawning of sharpnose puffers had occurred earlier according to Karl; this rare event had not been observed for 12 years. Many juvenile puffers (1 cm) were observed in shallower water 200-100 m. Lionfish were first observed at 100 m depth (time 14:39). Blue sponges were common on vertical faces of boulders; one green sponge was observed. Large (0.5 m?) glass sponges were commonly observed, as were vase sponges. We need names for sponges, as these are common faunal components.

DeepCASTFigure 8. Large II DivePrimnoidae 1 at 1900 Date: ft depth May (579 22, m, 2011 left) with Location: squat lobsters Luna and Beach Paramuriceidae (right) with Asteroschema sp. brittlestars at 1350 ft depth (411 m). 8

DeepCAST II Dive 1 Date: May 22, 2011 Location: Luna Beach

Dive profile for Dive 1 (timed out after 49 min)

Figure 9. A plot of depth and time showing a 46 minute surface tow and 3 minute descent

Water chemistry profile for Dive 1

Figure 10. A plot of salinity, dissolved oxygen, temperature and pH for Dive 1 at Luna Beach.

DeepCAST II Dive 1 Date: May 22, 2011 Location: Luna Beach

Dive 1 Chronology: Time Depth (ft) Temperature (°C) 1014 S 30 1019 200 28 540 21 800 18 900 17.5 1031 1000 16 1032 1100 15 1034 1240 13.5 1035 1300 13 1039 1400 Many small squid (<2 cm) & small hatchet fish. 1042 1485 12.5 No more squid observed 1046 1680 12 1049 1800 12 Juvenile frost fish observed 1900 12 1055 2000 11 1057 2090 11 1130 1850 10 Brittlestar with white & pink striped disk. 1200 1640 12 CHANGE TAPE tape 2. Parazoanthus occurred in patches. 1208 1500 Six gill shark (3-4 m) 1215 1500 Barren boulder 1237 1290 12 1309 1200 12 Changed video camera battery 1317 1190 12 Large boulder field 1344 1110 12 Left boulder field; large field of Paramuricea. Atop the boulder were lots of coral and eipifauna. 1403 990 Started observing light from surface. 850 Paramuricea with Asteroporpa annulata brittlestars 1430 380 25 1439 100

DeepCAST II Dive 2 Date: May 23, 2011 Location: Sueno del Mar

Descent coordinates: 16 18.287’ N, 86 36.646’ W Observers: Jerry Harasewych, Sebastian Troeng Left dock: 10:15 Begin Descent: 10:35 Return to dock: 15:10

Geology: Maximum depth of dive 2150 ft. (655 m) Steep basaltic wall with many large basaltic boulders, with occasional sand pockets. Massive basaltic boulder >100 ft high at 1180 ft, bottom more gently sloped [30-45°] at about 1000 ft (305 m), predominantly sand / sediment with occasional boulders. Travelled horizontally at about 1000 ft. Turned upslope to steeper [70 - 85°] slope with more rocks. Transitioned from basaltic rock to limestone at around 650-700 ft. Limestone wall nearly vertical, with overhangs, ledges and small crevices, until reef edge at about 200 ft (61 m).

Biology: Rocky bottom around 2000 ft (610 m) was only sparsely populated with small corals and large red sea stars, while sandy patches had occasional swimming holothurians, small, dark urchins, and anemones. White branched octocoral collected near 1980 ft (604 m). Lophelia pertusa was collected at 1920 ft (585 m). Dendrophyllia alternata was also collected. The number of glass sponges began to increase near 1800 ft (549 m), especially on vertical or near-vertical surfaces. Crinoids were present, including stalked, comatulid, and Holopus. Perotrochus midas was common between 1800 and 1200 ft (366 m). One Perotrochus cf. charlestonensis was collected at 1180 ft (360 m). At around 1100 ft (335 m), bottom became more gently sloped and primarily sand covered, with occasional rocky patches.

Octocorals on the rocks included large Primnoidae, smallish purple Paramuriceidae, white Muriceides colonies and many Bathyalcyon soft corals. Rocks also populated by sponges, Parazoanthus colonies and crustaceans [crabs and shrimp]. One clawed lobster seen around 1000 ft. Sandy areas had bluish, swimming holothurians, mostly single but occasionally in groups of 5 to 8. Sea urchins also occurred both singly and in groups. The wall became steeper, with a higher proportion of rocky bottom at depths of 1140-960 ft (350-292 m). Rocks usually covered with sponges, gorgonians, crinoids, crabs and rarely Perotrochus quoyanus. Near 600 ft (183 m), the wall became nearly vertical, with cracks, ravines, overhangs and caverns. Fauna quite sparse at the base, becoming denser, mostly sponges, by 400 ft (122 m). Numerous tube sponges by 300 ft (91 m). Entemnotrochus adansonianus collected at 260 ft (79 m).

Figure 11. An aggregation of Lophelia pertusa at 1920 feet (585 m) depth at Sueno del Mar.

DeepCAST II Dive 2 Date: May 23, 2011 Location: Sueno del Mar

Dive profile for Dive 2 (timed out after 45 min)

Figure 12. A plot of depth and time showing a 40 minute surface tow and 6 minute descent

Water chemistry profile for Dive 2

Figure 13. A plot of salinity, dissolved oxygen, temperature and pH for Dive 2 at Sueno del Mar.

DeepCAST II Dive 2 Date: May 23, 2011 Location: Sueno del Mar

Dive 2 Chronology: Time Depth (ft) Temperature (°C) Comment 10:35 0 30.5 10:43 400 25.0 10:46 500 22.0 10:48 680 20.0 10:52 1000 16.0 10:58 1420 12.5 11:02 1540 12.0 11:06 1980 11.5 11:17 2150 10.5 bottom 11:26 2000 11.5 11:35 1920 11.5 Lophelia 11:45 1780 12.0 midas 1 [large] 11:53 1680 12.0 11:57 1660 12.0 midas 2 [small] 12:07 1560 12.0 frogfish 12:11 1500 12.0 midas [not collected] 12:14 1480 12.0 CHANGE TAPE 12:21 1440 12.5 midas 3 & not collected 12:32 1300 13.0 blue holothurians 12:38 1180 14.0 giant basaltic boulder 12:48 1180 14.0 cf. charlestonensis 12:53 1160 14.0 midas 4 13:08 1140 14.0 midas [not collected] 13:16 1060 15.0 sea fan w/ crinoids 13:37 1000 15.5 boulder 13:49 1000 15.5 jack 13:58 980 15.5 sand bottom w/ rocks 14:10 960 15.5 quoyanus [not collected] 14:15 940 15.5 sponges 14:25 920 16.0 quoyanus 1 14:34 700 18.5 last basaltic rock 14:37 600 20.0 steep, limestone slope 14:40 460 22.5 limestone wall 14:43 360 25.5 3 jacks 14:47 260 27.0 adansonianus 1 14:50 200 29.5 reef flat edge 14:52 0 30.5 surfaced

DeepCAST II Dive 3 Date: May 23, 2011 Location: Half Moon Bay

Descent coordinates: Slightly north of buoy. Buoy is 16 18.434 N, 86 35.740 W Observers: Laney Thornton, Annie Thornton Left dock: 1900 Begin Descent: 1907 Return to Surface: 2054

No boat tow. Submersible left and returned under her own power.

Geology: Maximum depth of dive was 1530 ft. Biology: Scientists were not present on this dive. No biological description is provided.

Dive 3 Chronology:

Time Depth (ft) Temperature (°C) 1909 S 30 1912 150 29 1916 350 25 1918 500 22 1922 700 19 1928 1100 15 1935 1400 14 1939 1530 13 bottom camera on 1942 1530 13 coral rubble 1947 1510 13 sand 1948 1500 13 Limestone boulder 2002 1510 13 Collecting trap located 2005 1510 13 Trap picked up 2010 1430 13 Basalt 2014 1360 13 Limestone boulders 2020 1200 15 2021 1180 15 Water column 2026 1000 17 Sand 2035 700 18 coral rubble/old trap 2043 520 23 Going up vertical wall 2048 300 27 Going up wall 2054 S 33

DeepCAST II Dive 3 Date: May 23, 2011 Location: Half Moon Bay

Dive profile for Dive 3 (timed out after 45 min)

Figure 14. A plot of depth and time showing a 19 minute surface tow and a 26 minute descent

Water chemistry profile for Dive 3

Figure 15. A plot of salinity, dissolved oxygen, temperature and pH for Dive 3 at Half Moon Bay.

DeepCAST II Dive 4 Date: May 25, 2011 Location: Three Kings (AKR)

Descent coordinates: 16° 21.026’ N, 86° 34.348’ W Observers: Kate Lavelle, Annie Thornton Depart dock: 08:47 Begin descent: 09:10 Return to dock: 12:58 No CTD cast.

Geology: Maximum depth of dive 2160 ft (658 m). Many large basaltic boulders, fewer limestone boulders. Boulders recently moved by earthquake activity had fewer sessile organisms. No patterns in boulder distribution were observed. Soft sediment filled the space between boulders.

Biology: The most abundant corals on hard substrate were primnoids, which were observed at a maximum depth of 1800 ft (548 m). Other corals included sea whips, bamboo corals, Bathypathes, Corallium, Paramuricea, Dendrophyllia alternata, and Plumapathes. Juvenile galatheoid crabs were seen on bamboo corals and primnoids. Adult galatheoids were observed primarily on primnoids.

Several species of sponges were observed including glass and vase sponges, with the majority occurring on boulder faces or the tops of boulders. Two species of cidaroid urchins were seen on Corallium and Bathypathes corals. Elasipodid sea cucumbers, brittle stars, anemones, and crinoids were observed on soft sediment. The most conspicuous invertebrates were large brisingid seastars, which were often in close proximity to corals and on boulders. Epifauna of primnoid colonies included venus flytrap anemones, galatheoids, brittle stars, brisingids, and crinoids.

Two wahoo were seen above the sub while descending (680 ft, 207 m). Two large pyrosomes were seen in the water column at 1900 ft (579 m). An unknown red jellyfish was observed at 2000 ft (610 m, possibly Periphylla). Two midas slit shells were seen at 1940 ft (591 m). Trash bags, bottles, and plastics were observed in a sand channel beginning at 1880 ft (573 m). Five lionfish were seen at 250 ft (76 m).

Dive 4 chronology: Time Depth (ft) Comment 9:10 surface 9:11 200 temperature was not recorded on this dive 9:13 300 9:15 400 9:17 600 still see surface light 9:19 680 2 wahoo above sub 9:21 790 9:23 900 9:25 1000 no more light 9:27 1090 9:29 1250 9:31 1290 9:33 1450 9:35 1500

DeepCAST II Dive 4 Date: May 25, 2011 Location: Three Kings (AKR)

Dive 4 chronology (continued) Time Depth (ft) Comment 9:37 1600 2’ cutlass fish 9:39 1690 squid eggs and squid 9:41 1800 salps, siphonophores 9:43 1890 1.5’ pyrosome 9:45 2000 lights out, some bioluminescence 9:47 2100 9:49 2160 9:53 2120 siphonophore, steep incline 9:57 2100 9:59 2060 10:01 2000 lots of bare boulders, from recent earthquakes 10:03 2000 sea urchin, bathypathes 10:05 2000 red jelly, unknown 10:07 2000 clear jelly, maybe ctenophore 10:09 2000 very steep rockface 10:11 2000 large siphonophore 10:13 2000 10:15 2000 10:17 2000 huge pyrosome 10:19 2000 bamboo coral with juvenile crabs 10:21 2000 10:23 2000 10:25 2000 10:27 2000 starting to see more corals, steep 10:31 2060 goosefish, Plumapathes, primnoid, Corallium, bamboo 10:33 2000 10:35 2000 10:37 2000 10:39 2000 10:41 2000 10:43 2000 10:45 2000 10:49 bare Bathypathes, chirostylid nearby 10:53 2020 CHANGE TAPE tape 2 begins 10:57 2020 10:59 coasting 11:03 2020 sandy bottom, sand channels 11:05 2000 11:07 2000 some bamboo corals, green Plumapathes 11:11 2020 turned around back to where we saw coral

DeepCAST II Dive 4 Date: May 25, 2011 Location: Three Kings (AKR)

Dive 4 chronology (continued) Time Depth (ft) Comment 11:19 1940 two large midas 11:21 1880 trash bags, rusting metal 11:27 1800 11:29 1780 11:31 1640 steep slope 11:33 1600 12 C 11:35 1520 11:37 1500 can see ledge 70 ft above us 11:39 1490 cleared ledge 11:41 1460 large limestone rock 11:43 1400 small colonies Dendrophyllia 11:45 1380 11:47 1300 Paramuricea 11:49 1200 sponges cover boulders 11:51 1180 11:55 1100 11:59 1080 12:01 1040 garbage, cans, bottles 12:03 1040 garbage, four large unidentified fish 12:05 900 12:09 800 CHANGE TAPE tape 3, rope hanging at 800-850 ft 12:11 780 12:13 700 wall 250’ away 12:15 680 12:17 600 12:19 560 12:21 500 12:23 480 12:25 470 12:27 420 12:29 410 12:33 310 large Ellisellidae, presumably Nicella sp., five lionfish 12:35 250 current now moving opposite direction than current at 2000’ 12:37 200 visible thermocline 12:41 surfaced

DeepCAST II Dive 4 Date: May 25, 2011 Location: Three Kings (AKR)

Large sea fans in the families Primnoidae and Ellisellidae. Green lasers are 10 cm apart.

Figure 16. A large primnoid colony in the subfamily Calyptrophorinae at 1800 ft (548 m) depth.

Figure 17. A sea fan in family Ellisellidae with many brittlestars and trash (yellow) at 310 ft (94 m) depth.

DeepCAST II Dive 4 Date: May 25, 2011 Location: Three Kings (AKR)

Dive profile for Dive 4

Figure 18. A plot of depth and time showing a 35 minute descent with ~165 minutes of bottom time.

Water chemistry profile for Dive 4

Figure 19. A plot of salinity, dissolved oxygen, temperature and pH for the downcast of Dive 4 at Three Kings. An asterisk (*) indicates the depth (580 m) of Lophelia observed at this site on August 14, 2010.

DeepCAST II Dive 5 Date: May 26, 2011 Location: Lawson’s Rock

Descent coordinates: 16° 20.188’ N, 86° 34.267’ W Observers: Sebastian Troëng, Laney Thornton Depart dock: 09:05 Begin descent: 09:48 Back at surface: 14:27 Return to dock: 14:39

Geology: Maximum depth of dive 2,090 ft (637 m). Upon reaching sea bottom there were large basaltic boulders of volcanic origin that appeared striated. We observed a couple of cinderblocks with a rope stretching in a slight slope as far as we could see. Karl mentioned these may be from the Roatán Marine Park staff’s effort to install a buoy but underestimating the depth of the water and only having 1,500 ft of rope for the 2,000 ft depth. After the initial basaltic boulders there were large areas of soft bottom with occasional basaltic boulders and after the middle of the dive there were occasional calcium carbonate boulders including some very large ones. Two boulders in particular stood out, one that was separated by a small distance from the main basaltic boulders (observed at 11:13) and a second that was towards the end of the dive (observed 13:41), in front of the channel leading to Anthony’s Key Resort and that was covered in sea fans and epifauna. For much of the dive (the duration of the two first videotapes) we remained at depths exceeding 1,800 ft (549 m) followed by a slow ascent lasting two hours.

Biology: The amount of material in the water column varied considerably during the dive. Initially there were lots of materials in the water and the bottom fauna appeared more ample with soft bottom fauna dominated by lobsters, worms and brittlestars. Later during the dive the water was clearer with less material in the water column (and less rich bottom fauna) followed a little later by more material and finally by clearer water again. We observed a giant isopod when arriving at the sea bottom. Other noteworthy fauna includes a frostfish (photographed) and an unidentified shark (approx. 16 in. long) that we filmed but could not photograph (autofocus would not lock). We collected a midas slitshell and three coral genera (Dendrophyllia, Madrepora, Enallopsammia). During the ascent we observed large groups of urchins at 1,000 ft (305 m) and on a sandy slope at 200 ft (61 m) we observed numerous garden eels.

Figure 20. Sample collection of Dendrophyllia alternata from Lawson’s Rock.

DeepCAST II Dive 5 Date: May 26, 2011 Location: Lawson’s Rock

Dive 5 chronology: Time Depth (ft) Temp (°C) Comment 9:48 S 31 9:51 200 29 9:53 400 24 9:56 600 20.5 9:58 800 18 10:01 1000 16 10:04 1200 14 10:06 1400 13 10:10 1600 12.5 10:14 1800 12 10:17 2000 12 10:20 2090 11.5 bottom, volcanic rock, striation rock may be sedimentary 10:22 11.5 giant isopod 10:28 2040 11.5 volcanic rock, some with striation, black corals 10:30 2040 11.5 sandy bottom with lobster and cinderblocks with rope from Roatán Marine Park (according to Karl) 10:37 2060 11.5 volcanic rock with gorgonians and Madrepora 10:45 2040 11.5 another volcanic rock with gorgonians, black corals, Madrepora 10:53 2040 12 frostfish 10:58 2020 12 sandy bottom, lobsters, worms, brittlestars 11:13 1960 11.5 steep cliff to left with limestone slab to right, less fish life 11:23 1940 12 changed to tape 2 11:35 2040 12 sediment w/ occasional volcanic ridge, less material in water 11:47 2010 12 seafans with sea stars 12:02 1940 12 small overhang, more material in water column 12:08 1910 12 changed battery 12:20 1980 12 midas slitshell collected, volcanic rock 12:26 1930 12 Enallopsammia coral collected 12:27 1960 12 tube sponge photographed, changed to tape 3 12:37 1820 12 midas on sand, midas on rock 12:45 1680 12.5 ascending volcanic boulder, slope with corals and garbage 12:53 1560 12.5 green sponges, volcanic slope 12:58 1460 12.5 yellow coral collected 13:05 1400 12.5 Dendrophyllia collected, midas on coral 13:12 1340 13 midas on rock 13:18 1300 13.5 Dendrophyllia 13:25 1230 13.5 coral sampled, shelled mollusk on wall 13:41 1160 14 calcium carbonate rock 13:46 1100 14 rock covered in seafans 13:53 1020 14 back on sandy slope, numerous urchins together 14:01 820 18 sandy slope with garbage 14:06 680 19.5 airplane part in sand 14:11 500 22 going up steep calcium carbonate drop-off 14:14 400 24 14:21 200 30 garden eels 14:27 surface 31 14:39 surface 33 dock

DeepCAST II Dive 5 Date: May 26, 2011 Location: Lawson’s Rock

Large sea fans in the families Primnoidae and Paramuriceidae. Green lasers are 10 cm apart.

Figure 21. A large sea fan in family Primnoidae, with many squat lobsters at 1100 ft/335 m depth.

Figure 22. Sea fans in family Paramuriceidae with Asteroschema sp. brittlestars at 1230 ft/374 m depth.

DeepCAST II Dive 5 Date: May 26, 2011 Location: Lawson’s Rock

Dive profile for Dive 5 (timed out after 45 min)

Figure 23. A plot showing a 15 min. surface tow, a 26 min. descent, and 90 min. of along-track data.

Water chemistry profile for Dive 5

Figure 24. A plot of salinity, dissolved oxygen, temperature and pH for Dive 5 at Lawson’s Rock. An asterisk (*) indicates observed depth of Lophelia at Sueno del Mar on May 23, 2011 near 585 m depth.

DeepCAST II Dive 6 Date: May 27, 2011 Location: Old Loafers

Descent coordinates: 16° 17.924’ N, 86° 36.292’ W Observers: Tom Shirley, Jerry Harasewych Left dock: 1415 Begin Descent: 1448 Return to Surface: 1833

Geology: Maximum depth of dive was 1220 ft (372 m). Sub release point from boat was too shallow, only 700 ft (213 m) deep and sub had to drive offshore, near bottom, to find deeper depths. Many large boulders (limestone?) were interspersed with soft sediment. Many boulders were sediment-covered and supported few epifauna. At depths of about 500 ft (152 m), the slope became nearly vertical, a limestone wall that was deeply eroded [like a meteorite] in areas, with the top of the wall at around 200 ft (61 m).

Biology: Soft substrate was occupied by sea urchins, a few sponges, and elasipodid sea cucumbers. One cat shark was observed, perhaps 1 m in total length; sub traveled directly over it and we had a good view through the bottom view port. Octocorals were relatively uncommon, mainly the purple/grey sea fans that I refer to as Paramuricea, and always on hard substrate, usually on tops of boulders. Crinoids were among the most conspicuous animals on this dive, occurring on a variety of different substrates: boulder, sponges, and on the few octocorals.

One site near the beginning of the dive had a natural arch and was covered with sea lilies, more than Karl had previously observed. We named this site “Sea Lily City.” Brisingid seastars (Novodinia antillensis) occurred on many substrates, but were not as common as on previous dives. Elasipodid sea cucumbers were usually solitary, often several meters from other cucumbers on soft sediment. A small (20-39) school of shrimp was observed on the bottom near an anemone. One small aggregation of sea urchins, about 10 sea urchins, occurred on soft sediment in a group, near the beginning of the dive. Two species of slit shells were observed – one on tops of boulders, the other on near vertical walls. A new species of turrid shell was collected.

A bamboo coral (Lepidisis) occurred in solitary stalks commonly, and was more common in shallower depths. Galatheoid crabs occurred on some of the Lepidisis. A new, small brittlestar species was observed on what appeared to be a yellow Paramuricea (but with slightly larger branches) in much shallower water (450 ft, 137 m). These brittlestars blended in almost perfectly with the coral and were difficult to distinguish. Many juvenile sharpnose puffers (1-2 cm in length) were observed in shallower waters (400-200 ft). Three lionfish were first observed at 400 ft (122 m) depth, but then lionfish were observed continuously to the surface. Blue sponges were common on vertical faces of some boulders.

Figure 25. Corals at Old Loafers included sea fans in family Paramuriceidae (1200 ft/366 m, left) and large thickets of the scleractinian coral Madracis myriaster (400 ft/122 m, right). Green lasers are 10 cm apart.

DeepCAST II Dive 6 Date: May 27, 2011 Location: Old Loafers

Dive 6 chronology: Time Depth (ft) Temperature (°C) 1420 S 33 1435 S 32 1445 S 32 1448 40 30 Beginning dive 1450 100 30 1451 200 29.5 1452 300 28 1453 400 24 1455 500 23 1456 600 22 1458 700 20 Near bottom, heading offshore to deeper water 1501 800 19 1504 900 18 1506 1000 16 Can see bottom 1508 1100 15.5 Temperatures are almost identical to Dive 1 (at same depths) 1509 1150 15 Soft sediment bottom – fine sand/silt 1512 1220 14 Lots of lebenspuren visible: tracks, ruts, mounds, burrows, depressions 1520 1240 13 Moving across sand bottom, around large boulder. 1525 1200 13 Several small, silvery fish hanging upside down under Paramuricea 1535 1180 14 Limestone rx with galatheoid 1542 1180 14 Silty bottom with limestone rx 1547 1100 “Sea Lilly City” on large boulder with arch 1555 1100 15 carrier shell, Parazoanthus 1602 1100 14 Black Rock (Karl’s name for site) – Large Antipathes sp black coral 1610 1100 14 Level bottom, pits and mounds 1611 1100 14 Bathypathes (black coral resembling a feather-shape) w/ large galatheoid 1615 1100 16 CHANGED TAPE; 1625 1110 16 Collected slit shell 1638 1060 Collected turrid shell 1643 1000 16 Large boulders, lots of sediment 1645 900 17 Large boulders, lots of sediment 1654 890 Cat shark; squat lobster 1700 840 Switched camera to AC power; battery ran out; lizard fish 1705 800 18 Calliostoma sp. collected; 1.5 inches (?) in diameter 1715 740 Large vase sponges on large boulders; sediment covered 1722 650 18 CHANGED TAPE; Engine block & 100 ft. of line; little coral observed 1730 500 21 Drum fish (like a high hat); steep slope 1735 400 24 Steep slope; French angel; short bigeye; Madracis myriaster 1738 400-380 3 lionfish; 1745 390 One very large lionfish 1757 390 26 Steep wall, barren 1801 320 27 Slit shell collected (Entemnotrochus adansonianus); 2 large lionfish 1810 400 26 CHANGED TAPE; tape 4; collected Madracis; lots of Lionfish, ~14” 1815 380 26 Collected more Madracis 1825 300 28 Collected another slit shell (Entemnotrochus adansonianus); another slit shell was within a few cm; on steep wall, near sponge 1833 Surfaced

DeepCAST II Dive 6 Date: May 27, 2011 Location: Old Loafers

Dive profile for Dive 6 (complete profile)

Figure 26. A plot showing a 20 min. surface tow, a 15 min. descent, and 190 min. of along-track data.

Water chemistry profile for Dive 6

Figure 27. A plot of salinity, dissolved oxygen, and temperature during the 15 min. descent at Old Loafers.

Conclusion:

The DeepCAST II Expedition represented this research team’s second scientific foray onto the deep-reefs of Roatán, Honduras. Research methods and techniques improved substantially since the first three day expedition in May 2010 (now referred to as DeepCAST I). Notable improvements include exterior mounts for cameras and lasers, along-track water chemistry from NOAA’s Seabird CTD-O, and two new methods of sample collecting, the long-pole net and the baited trap. The primary significance of the research was enhanced knowledge of the deep-sea diversity present in the deep reaches of the MAR, and new baseline information on the environment, health and condition of deep-sea corals off Roatán, Honduras.

Biological sampling was identified as the first research priority after DeepCAST I. We were highly successful in this regard. The long-pole net retrieved 19 of 20 specimens (see Appendix 1 and 2), including five species of constructional scleractinians, four species of slit shells, one precious octocoral (genus Corallium), and one new species of turrid shell. Photo samples documented many more species (Appendix 3). We collected more photographic evidence of coral depredation, but have yet to identify the predator. Slit shells and cidaroid pencil urchins are currently the primary suspects, so future sampling efforts will target these animals.

The research team has grown to include new partners (e.g., Dr. Jerry Harasewych from Smithsonian NMNH), and new members from established partners (e.g., master’s student Kate Lavelle from TAMU- CC, Dr. Sebastian Troeng from Conservation International). Results of our fieldwork have also opened the door to future collaborations with research partners in other taxa, like fishes and crinoids.

The next steps for field work are: 1) to improve the duration and performance of the along-track water chemistry profiler so every photo-transect can be cross-referenced to environmental parameters (e.g., depth and temperature), 2) to collect more samples of corals and associated invertebrates, and 3) to explore new habitats for better quantification of alpha and beta diversity on Roatán’s deep-reefs.

Long term goals include: 1) high-resolution multibeam mapping of the Roatán Escarpment, 2) baseline characterization of aragonite saturation to 1000 m depth, and 3) more intensive biological sampling for population genetics, to understand patterns of connectivity between deep-reef assemblages along the Meso-American Reef and deep-reef assemblages in the adjacent Gulf of Mexico.

Acknowledgements:

This research would not have been possible without the generous support of Dr. Jeff Hyland at NOAA’s Coastal Center for Environmental Health and Biomolecular Research; and Drs. Steven Cairns and Jerry Harasewych at Smithsonian National Museum of Natural History. Conservation International’s (CI) Sojourns Program contributed greatly to our success. Rod Mast, Levi Norton, and Karl Egloff were instrumental to our success, along with Laney Thornton and Anne Chapman-Thornton. Dr. Sebastian Troeng of CI contributed important notes and insights. Schmidt Ocean Institute and Dr. Fred Boltz of CI helped catalyze the research through their participation in DeepCAST I.

Karl Stanley’s efforts, ideas, and generosity were largely responsible for much of what we were able to accomplish during the expeditions. Research Fellowship funds from Schmidt Ocean Institute provided new tools, including new cameras and lasers. Dr. Steve W. Ross of University of North Carolina- Wilmington provided fish identifications. Dr. Cairns identified the scleractinians. Andrés Alegría, Coordinador de Monitoreo Biológico, Instituto Nacional de Conservación y Desarrollo Forestal, Áreas Protegidas y Vida Silvestre (ICF), Honduras kindly assisted with sample processing, as did Dr. Steve Box of Utila Ecology. Thanks to everyone involved for their generous support.

Appendix 1. DeepCAST II Biological Sample Inventory

The following specimens were collected as part of the Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) Expedition in Roatán, Honduras, May 21-28, 2011. Specimens were collected under the authority of Resolucion DE-MP-074-2011 from Instituto Nacional de Conservacion y Desarollo Forestal, Areas Protegidas y Vida Silvestre.

Scleractinia are regulated under the Convention for International Trade of Endangered Species (CITES). Other marine invertebrates listed in the collecting permit are exempt from CITES.

Specimens collected that are regulated by CITES:

IDA-11-01 Dendrophyllia alternata – 1 tube in ethanol, 1 bag dry IDA-11-04 Lophelia pertusa – 1 tube in ethanol, 1 bag dry IDA-11-05 Madrepora oculata - 1 tube in ethanol, 1 bag dry IDA-11-06 Enallopsammia rostrata - 1 tube in ethanol, 1 bag dry IDA-11-07 Dendrophyllia alternata – 1 tube in ethanol IDA-11-10 Madracis myriaster – 1 bag dry

Specimens that are not regulated under CITES:

Miscellaneous Invertebrate samples: IDA-11-03 Tube 10 - Octocoral, Corallidae, white IDA-11-09 Tube 11 - Sponge, white, thin IDA-11-02 Tube 12 – Bryozoa IDA-11-08 Tube 13 – Holopus crinoids IDA-11-11 2 specimens of lithodid crab Paralomis cubensis; a female and larger male

Mollusk samples [FM = foot muscle] Tube 01 = Perotrochus cf. chalestonensis FM ex living animal. Tube 02 = Perotrochus midas FM smallest specimen ex living animal. Tube 03 = Perotrochus midas FM second smallest specimen ex living animal. Tube 04 = Perotrochus midas FM medium sized, broken specimen ex living animal. Tube 05 = Perotrochus midas FM largest specimen ex living animal. Tube 06 = Perotrochus quoyanus FM ex living animal. Tube 07 = Perotrochus cf. chalestonensis FM ex tissue provided by Karl Stanley Tube 08 = Entemnotrochus adansonianus Dive 2 FM ex living animal. Tube 09 = Perotrochus quoyanus FM from tissue sample provided by Karl Stanley.

As per the collecting permit application, up to three samples per species will be deposited into the collections of the National Museum of Natural History, Smithsonian Institution, where the collecting information, catalog number and images of specimens will be available on line. For those samples that result in successful sequencing, the sequence data will be made available on GenBank. Specimens additional to those permitted to be retained at SI will be returned to the Museo Nacional in Honduras.

Appendix 2. DeepCAST II Biological Sample Photo Inventory

Figure A1. Sample IDA-11-01 Dendrophyllia alternata

Figure A2. Sample IDA-11-04 Lophelia pertusa